C. elegans gene mlt-11 is a Kunitz family putative protease inhibitor required for proper cuticle formation and molting. Using tissue-specific RNA interference we tested in which tissues mlt-11 activity was critical for molting and developmental progression. mlt-11 knockdown using a seam cell-enriched RNAi strain produced similar phenotypes as knockdown throughout animals. In contrast, hypodermis-enriched knockdown caused no detectable phenotype. Together, our findings identify seam cells as a critical site of mlt-11 function and highlight an unexpectedly central role for this specialized epithelial cell type in regulating molting and developmental progression.
","acknowledgements":"We thank Zoe Johnson, Javier Hernandez Lopez, Zoie Reyna, Emma Cadena, Valarie Hallin, and Olivia Vedar for research support. Some strains were provided by the Caenorhabditis Genetics Center, which is funded by the NIH Office of Research Infrastructure Programs (P40 OD010440). Wormbase was used in the design and execution of experiments. Wormbase and the Alliance of Genome Resources were used in the design and execution of experiments.
","authors":[{"affiliations":["Department of Molecular, Cell, and Developmental Biology, University of California – Santa Cruz, Santa Cruz, CA 95064"],"departments":[""],"credit":["investigation","methodology","writing_reviewEditing"],"email":"jragle@ucsc.edu","firstName":"James Matthew","lastName":"Ragle","submittingAuthor":false,"correspondingAuthor":null,"equalContribution":null,"WBId":"","orcid":"0000-0002-6626-2615"},{"affiliations":["Department of Molecular, Cell, and Developmental Biology, University of California – Santa Cruz, Santa Cruz, CA 95064"],"departments":[""],"credit":["formalAnalysis","fundingAcquisition","investigation","methodology","supervision","validation","visualization","writing_originalDraft"],"email":"jward2@ucsc.edu","firstName":"Jordan D.","lastName":"Ward","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":null,"WBId":"","orcid":"0000-0001-9870-8936"}],"awards":[],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":null},"extendedData":[],"funding":"This work was funded by the National Institutes of Health (NIH) National Institute of General Medical Sciences (NIGMS) (R01GM138701, and R35GM158317) to J.D.W.
","image":{"url":"https://portal.micropublication.org/uploads/5fca52711c3166238cb30a6745c2d5d9.png"},"imageCaption":"Tissue-specific RNAi strains used. A timed egg lay of animals of the indicated genotype were performed on control or mlt-11 (RNAi) plates and phenotypes were scored three days later. (B) Representative images of animals of the indicated genotype grown on control or mlt-11 RNAi. The red box highlights conditions that produced smaller larvae with molting defects. (C) Developmental delay was scored in the indicated strains grown on control or mlt-11 RNAi plates and classified as a failure to reach adulthood after 72 hours of growth. (D) Molting defects were scored in the indicated strains on control or mlt-11 RNAi. Scored defects included animals dragging cuticles, ecdysis failure and cuticle corsets. Tissue-specific RNAi data is from two independent replicates and the number of animals scored is in (A).
","imageTitle":"mlt-11 is necessary in seam cells for molting and larval development
","methods":"RNAi Knockdown
RNA interference experiments were performed as in Johnson et al. (2023). Control RNAi used an empty L4440. The mlt-11 (RNAi) vector was streaked from the Ahringer library (Kamath et al., 2003).
","reagents":"Strain name | Genotype | Source |
Bristol isolate, standard lab wild type | CGC | |
jsTi1493 {mosL loxP [wrdSi72(SCMp::pes-10delta::rde-1 CDS+3'UTR)] FRT3::mosR} IV ; rde-1(ne300) V | Ward lab (Johnson et al., 2023) | |
jsTi1493 {mosL loxP [wrdSi97(suro-1p::rde-1 CDS+3'UTR)] FRT3::mosR} IV ; rde-1 (ne300) V | Ward lab (Johnson et al., 2023) | |
rde-1(ne300) V ; neIs9(neIs9 [myo-3::HA::RDE-1 + rol-6(su1006)]) X | ||
Molting is an essential developmental process in nematodes, involving periodic synthesis of a new collagen-rich apical extracellular matrix (cuticle) and the shedding of the old cuticle. Understanding the mechanisms underlying molting can provide insight into oscillatory biological timers, epithelial biology, and is a source of potential targets to develop anthelminthics to control parasitic nematodes of medical and agricultural importance. Proteases and protease inhibitors play important and poorly understood roles in remodeling of the C. elegans aECM during molting.
mlt-11 is a putative Kunitz family protease inhibitor that plays a critical role in C.elegans cuticle synthesis and molting (Frand et al., 2005; Ragle et al., 2026). It is a complex protein with 10 Kunitz domains and depending on which of these domains are deleted, phenotypes such as left rollers, right rollers, microblistering, or embryonic lethality (Ragle et al., 2026). Reduction of mlt-11 function results in aberrant protein localization in the aECM over both seam and hypodermal cells, and all three layers of the adult cuticle were affected (basal, medial, and cortical layers) (Ragle et al., 2026). Given that mlt-11 is expressed in both seam cells and in the hypodermal syncytium (Frand et al., 2005; Ragle et al., 2026), we turned to a tissue-specific RNAi approach to test where mlt-11 activity was needed to promote molting (Fig. 1A). mlt-11 knockdown in a tissue-specific RNAi strain (JDW371) that restricts knockdown to seam, hypodermal, and intestinal cells (Johnson et al., 2023), phenocopied mlt-11 RNAi in wildtype animals with respect to developmental delay and molting defects (Fig. 1B-D). mlt-11 RNAi in a hypodermal and seam cell-specific RNAi strain (QK52), produced less penetrant developmental delay and molting defects (Fig. 1B-D). Notably, mlt-11 RNAi in a hypodermal-specific RNAi strain (JDW510),produced no developmental delay or molting defects, suggesting mlt-11 activity is necessary in seam cells and perhaps, in a limited capacity, in other hypodermal cells (Fig. 1B-D).
Despite robust hypodermal expression and localization (Frand et al., 2005; Ragle et al., 2026), our tissue-specific RNAi suggests that the seam cells are an important site of mlt-11 expression (Fig. 1). We consistently saw the most severe RNAi phenotypes in tissue-specific RNAi strains using a seam cell enriched promoter and no phenotypes from a hypodermal/intestinal-specific RNAi strain (Fig. 6). Consistent with these data, we previously observed aberrant expression of the markers DPY-7::GFP (furrows), DPY-10::mScarlet (furrows), ROL-6::mNG (basal layer collagen), CUT-2::mNG (cortical layer cuticlin), and COL-19::mNG (basal layer collagen) over the seam cells as well as alae gaps following mlt-11 inactivation (Ragle et al., 2026). The medial layer strut collagen BLI-1 is normally excluded from the aECM over the seam cells but invades this area following mlt-11 knockdown (Ragle et al., 2026). These data were reminiscent of our work on NHR-23 as depletion of this factor causes aberrant aECM formation over the seam cells and nhr-23 activity also was necessary in the seam cells for developmental progression (Johnson et al., 2023). Seam cells are responsible for the production of alae (Singh & Sulston, 1978), so it is unclear why inactivation of nhr-23 or mlt-11 in the seam cells has such a potent effect on developmental progression and molting given that the hypodermis secretes the bulk of the cuticle. Defining the relative contribution of each epithelial cell type to cuticle function and developmental progression is an important future direction.
","references":[{"reference":"Barkoulas M, van Zon JS, Milloz J, van Oudenaarden A, Félix MA. 2013. Robustness and epistasis in the C. elegans vulval signaling network revealed by pathway dosage modulation. Dev Cell 24(1): 64-75.
","pubmedId":"23328399","doi":""},{"reference":"Espelt MV, Estevez AY, Yin X, Strange K. 2005. Oscillatory Ca2+ signaling in the isolated Caenorhabditis elegans intestine: role of the inositol-1,4,5-trisphosphate receptor and phospholipases C beta and gamma. J Gen Physiol 126(4): 379-92.
","pubmedId":"16186564","doi":""},{"reference":"Frand AR, Russel S, Ruvkun G. 2005. Functional genomic analysis of C. elegans molting. PLoS Biol 3(10): e312.
","pubmedId":"16122351","doi":""},{"reference":"Johnson LC, Vo AA, Clancy JC, Myles KM, Pooranachithra M, Aguilera J, et al., Ward JD. 2023. NHR-23 activity is necessary for C. elegans developmental progression and apical extracellular matrix structure and function. Development 150(10): 10.1242/dev.201085.
","pubmedId":"37129010","doi":""},{"reference":"Kamath RS, Fraser AG, Dong Y, Poulin G, Durbin R, Gotta M, et al., Ahringer J. 2003. Systematic functional analysis of the Caenorhabditis elegans genome using RNAi. Nature 421(6920): 231-7.
","pubmedId":"12529635","doi":""},{"reference":"Melo JA, Ruvkun G. 2012. Inactivation of Conserved C. elegans Genes Engages Pathogen- and Xenobiotic-Associated Defenses. Cell 149: 452-466.
","pubmedId":"","doi":"10.1016/j.cell.2012.02.050"},{"reference":"Ragle JM, Turzo A, Jackson A, Vo AA, Pham VT, Daly K, et al., Ward. 2025. Roles and regulation of the Kunitz domain protein MLT-11 during\n C. elegans\n cuticle synthesis and molting. GENETICS 232: 10.1093/genetics/iyaf265.
","pubmedId":"","doi":"10.1093/genetics/iyaf265"},{"reference":"Singh, R. N., & Sulston, J. E. (1978). Some observations on molting in C. elegans. Nematologica, 24, 63–71.
","pubmedId":"","doi":""},{"reference":"Watts JS, Harrison HF, Omi S, Guenthers Q, Dalelio J, Pujol N, Watts JL. 2020. New Strains for Tissue-Specific RNAi Studies in Caenorhabditis elegans. G3 (Bethesda) 10(11): 4167-4176.
","pubmedId":"32943454","doi":""}],"title":"C. elegans mlt-11 activity is necessary in seam cells for molting and larval development
","reviews":[{"reviewer":{"displayName":"Xiaofei Bai"},"openAcknowledgement":false,"status":{"submitted":true}}],"curatorReviews":[{"curator":{"displayName":"Gary Craig Schindelman"},"openAcknowledgement":false,"submitted":null},{"curator":{"displayName":"KJ Yook"},"openAcknowledgement":false,"submitted":null}]},{"id":"3cec5431-c0ad-4323-b58b-c24e9f41a2ba","decision":"accept","abstract":"C. elegans gene mlt-11 is a Kunitz family putative protease inhibitor required for proper cuticle formation and molting. Using tissue-specific RNA interference we tested in which tissues mlt-11 activity was critical for molting and developmental progression. mlt-11 knockdown using a seam cell-enriched RNAi strain produced similar phenotypes as knockdown throughout animals. In contrast, hypodermis-enriched knockdown caused no detectable phenotype. Together, our findings identify seam cells as a critical site of mlt-11 function and highlight an unexpectedly central role for this specialized epithelial cell type in regulating molting and developmental progression.
","acknowledgements":"We thank Zoe Johnson, Javier Hernandez Lopez, Zoie Reyna, Emma Cadena, Valarie Hallin, and Olivia Vedar for research support. Some strains were provided by the Caenorhabditis Genetics Center, which is funded by the NIH Office of Research Infrastructure Programs (P40 OD010440). Wormbase was used in the design and execution of experiments. Wormbase and the Alliance of Genome Resources were used in the design and execution of experiments.
","authors":[{"affiliations":["Department of Molecular, Cell, and Developmental Biology, University of California – Santa Cruz, Santa Cruz, CA 95064"],"departments":[""],"credit":["investigation","methodology","writing_reviewEditing"],"email":"jragle@ucsc.edu","firstName":"James Matthew","lastName":"Ragle","submittingAuthor":false,"correspondingAuthor":null,"equalContribution":null,"WBId":"","orcid":"0000-0002-6626-2615"},{"affiliations":["Department of Molecular, Cell, and Developmental Biology, University of California – Santa Cruz, Santa Cruz, CA 95064"],"departments":[""],"credit":["formalAnalysis","fundingAcquisition","investigation","methodology","supervision","validation","visualization","writing_originalDraft"],"email":"jward2@ucsc.edu","firstName":"Jordan D.","lastName":"Ward","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":null,"WBId":"","orcid":"0000-0001-9870-8936"}],"awards":[],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":null},"extendedData":[],"funding":"This work was funded by the National Institutes of Health (NIH) National Institute of General Medical Sciences (NIGMS) (R01GM138701, and R35GM158317) to J.D.W.
","image":{"url":"https://portal.micropublication.org/uploads/5fca52711c3166238cb30a6745c2d5d9.png"},"imageCaption":"Tissue-specific RNAi strains used. A timed egg lay of animals of the indicated genotype were performed on control or mlt-11 (RNAi) plates and phenotypes were scored three days later. (B) Representative images of animals of the indicated genotype grown on control or mlt-11 RNAi. The red box highlights conditions that produced smaller larvae with molting defects. (C) Developmental delay was scored in the indicated strains grown on control or mlt-11 RNAi plates and classified as a failure to reach adulthood after 72 hours of growth. (D) Molting defects were scored in the indicated strains on control or mlt-11 RNAi. Scored defects included animals dragging cuticles, ecdysis failure and cuticle corsets. Tissue-specific RNAi data is from two independent replicates and the number of animals scored is in (A).
","imageTitle":"mlt-11 is necessary in seam cells for molting and larval development
","methods":"RNAi Knockdown
RNA interference experiments were performed as in Johnson et al. (2023). Control RNAi used an empty L4440. The mlt-11 (RNAi) vector was streaked from the Ahringer library (Kamath et al., 2003).
","reagents":"Strain name | Genotype | Source |
Bristol isolate, standard lab wild type | CGC | |
jsTi1493 {mosL loxP [wrdSi72(SCMp::pes-10delta::rde-1 CDS+3'UTR)] FRT3::mosR} IV ; rde-1(ne300) V | Ward lab (Johnson et al., 2023) | |
jsTi1493 {mosL loxP [wrdSi97(suro-1p::rde-1 CDS+3'UTR)] FRT3::mosR} IV ; rde-1 (ne300) V | Ward lab (Johnson et al., 2023) | |
rde-1(ne300) V ; neIs9(neIs9 [myo-3::HA::RDE-1 + rol-6(su1006)]) X | ||
Molting is an essential developmental process in nematodes, involving periodic synthesis of a new collagen-rich apical extracellular matrix (cuticle) and the shedding of the old cuticle. Understanding the mechanisms underlying molting can provide insight into oscillatory biological timers, epithelial biology, and is a source of potential targets to develop anthelminthics to control parasitic nematodes of medical and agricultural importance. Proteases and protease inhibitors play important and poorly understood roles in remodeling of the C. elegans aECM during molting.
mlt-11 is a putative Kunitz family protease inhibitor that plays a critical role in C.elegans cuticle synthesis and molting (Frand et al., 2005; Ragle et al., 2026). It is a complex protein with 10 Kunitz domains and depending on which of these domains are deleted, phenotypes such as left rollers, right rollers, microblistering, or embryonic lethality (Ragle et al., 2026). Reduction of mlt-11 function results in aberrant protein localization in the aECM over both seam and hypodermal cells, and all three layers of the adult cuticle were affected (basal, medial, and cortical layers) (Ragle et al., 2026). Given that mlt-11 is expressed in both seam cells and in the hypodermal syncytium (Frand et al., 2005; Ragle et al., 2026), we turned to a tissue-specific RNAi approach to test where mlt-11 activity was needed to promote molting (Fig. 1A). mlt-11 knockdown in a tissue-specific RNAi strain (JDW371) that restricts knockdown to seam, hypodermal, and intestinal cells (Johnson et al., 2023), phenocopied mlt-11 RNAi in wild-type animals with respect to developmental delay and molting defects (Fig. 1B-D). mlt-11 RNAi in a hypodermal and seam cell-specific RNAi strain (QK52), produced less penetrant developmental delay and molting defects (Fig. 1B-D). Notably, mlt-11 RNAi in a hypodermal-specific RNAi strain (JDW510),produced no developmental delay or molting defects, suggesting mlt-11 activity is necessary in seam cells and perhaps, in a limited capacity, in other hypodermal cells (Fig. 1B-D).
Despite robust hypodermal expression and localization (Frand et al., 2005; Ragle et al., 2026), our tissue-specific RNAi suggests that the seam cells are an important site of mlt-11 expression (Fig. 1). We consistently saw the most severe RNAi phenotypes in tissue-specific RNAi strains using a seam cell enriched promoter and no phenotypes from a hypodermal/intestinal-specific RNAi strain (Fig. 1B-D). Consistent with these data, we previously observed aberrant expression of the markers DPY-7::GFP (furrows), DPY-10::mScarlet (furrows), ROL-6::mNG (basal layer collagen), CUT-2::mNG (cortical layer cuticlin), and COL-19::mNG (basal layer collagen) over the seam cells as well as alae gaps following mlt-11 inactivation (Ragle et al., 2026). The medial layer strut collagen BLI-1 is normally excluded from the aECM over the seam cells but invades this area following mlt-11 knockdown (Ragle et al., 2026). These data were reminiscent of our work on NHR-23 as depletion of this factor causes aberrant aECM formation over the seam cells and nhr-23 activity also was necessary in the seam cells for developmental progression (Johnson et al., 2023). Seam cells are responsible for the production of alae (Singh & Sulston, 1978), so it is unclear why inactivation of nhr-23 or mlt-11 in the seam cells has such a potent effect on developmental progression and molting given that the hypodermis secretes the bulk of the cuticle. mlt-11 inactivation has been shown to cause reduced body length and width, impaired cuticle barrier function, and aberrant aECM protein localization (Ragle et al., 2026). Defining the tissue-specific requirements for mlt-11 in preventing these defects will be an important avenue for future investigation.”
","references":[{"reference":"Barkoulas M, van Zon JS, Milloz J, van Oudenaarden A, Félix MA. 2013. Robustness and epistasis in the C. elegans vulval signaling network revealed by pathway dosage modulation. Dev Cell 24(1): 64-75.
","pubmedId":"23328399","doi":""},{"reference":"Espelt MV, Estevez AY, Yin X, Strange K. 2005. Oscillatory Ca2+ signaling in the isolated Caenorhabditis elegans intestine: role of the inositol-1,4,5-trisphosphate receptor and phospholipases C beta and gamma. J Gen Physiol 126(4): 379-92.
","pubmedId":"16186564","doi":""},{"reference":"Frand AR, Russel S, Ruvkun G. 2005. Functional genomic analysis of C. elegans molting. PLoS Biol 3(10): e312.
","pubmedId":"16122351","doi":""},{"reference":"Johnson LC, Vo AA, Clancy JC, Myles KM, Pooranachithra M, Aguilera J, et al., Ward JD. 2023. NHR-23 activity is necessary for C. elegans developmental progression and apical extracellular matrix structure and function. Development 150(10): 10.1242/dev.201085.
","pubmedId":"37129010","doi":""},{"reference":"Kamath RS, Fraser AG, Dong Y, Poulin G, Durbin R, Gotta M, et al., Ahringer J. 2003. Systematic functional analysis of the Caenorhabditis elegans genome using RNAi. Nature 421(6920): 231-7.
","pubmedId":"12529635","doi":""},{"reference":"Melo JA, Ruvkun G. 2012. Inactivation of Conserved C. elegans Genes Engages Pathogen- and Xenobiotic-Associated Defenses. Cell 149: 452-466.
","pubmedId":"","doi":"10.1016/j.cell.2012.02.050"},{"reference":"Ragle JM, Turzo A, Jackson A, Vo AA, Pham VT, Daly K, et al., Ward. 2025. Roles and regulation of the Kunitz domain protein MLT-11 during\n C. elegans\n cuticle synthesis and molting. GENETICS 232: 10.1093/genetics/iyaf265.
","pubmedId":"","doi":"10.1093/genetics/iyaf265"},{"reference":"Singh, R. N., & Sulston, J. E. (1978). Some observations on molting in C. elegans. Nematologica, 24, 63–71.
","pubmedId":"","doi":""},{"reference":"Watts JS, Harrison HF, Omi S, Guenthers Q, Dalelio J, Pujol N, Watts JL. 2020. New Strains for Tissue-Specific RNAi Studies in Caenorhabditis elegans. G3 (Bethesda) 10(11): 4167-4176.
","pubmedId":"32943454","doi":""}],"title":"C. elegans mlt-11 activity is necessary in seam cells for molting and larval development
","reviews":[],"curatorReviews":[{"curator":{"displayName":"Gary Craig Schindelman"},"openAcknowledgement":false,"submitted":"1778776814082"},{"curator":{"displayName":"KJ Yook"},"openAcknowledgement":false,"submitted":null}]},{"id":"69f6306a-5904-4274-8bd5-8c617e5cdfcd","decision":"publish","abstract":"C. elegans gene mlt-11 is a Kunitz family putative protease inhibitor required for proper cuticle formation and molting. Using tissue-specific RNA interference we tested in which tissues mlt-11 activity was critical for molting and developmental progression. mlt-11 knockdown using a seam cell-enriched RNAi strain produced similar phenotypes as knockdown throughout animals. In contrast, hypodermis-enriched knockdown caused no detectable phenotype. Together, our findings identify seam cells as a critical site of mlt-11 function and highlight an unexpectedly central role for this specialized epithelial cell type in regulating molting and developmental progression.
","acknowledgements":"We thank Zoe Johnson, Javier Hernandez Lopez, Zoie Reyna, Emma Cadena, Valarie Hallin, and Olivia Vedar for research support. Some strains were provided by the Caenorhabditis Genetics Center, which is funded by the NIH Office of Research Infrastructure Programs (P40 OD010440). Wormbase was used in the design and execution of experiments. Wormbase and the Alliance of Genome Resources were used in the design and execution of experiments.
","authors":[{"affiliations":["Department of Molecular, Cell, and Developmental Biology, University of California – Santa Cruz, Santa Cruz, CA 95064"],"departments":[""],"credit":["investigation","methodology","writing_reviewEditing"],"email":"jragle@ucsc.edu","firstName":"James Matthew","lastName":"Ragle","submittingAuthor":false,"correspondingAuthor":null,"equalContribution":null,"WBId":"","orcid":"0000-0002-6626-2615"},{"affiliations":["Department of Molecular, Cell, and Developmental Biology, University of California – Santa Cruz, Santa Cruz, CA 95064"],"departments":[""],"credit":["formalAnalysis","fundingAcquisition","investigation","methodology","supervision","validation","visualization","writing_originalDraft"],"email":"jward2@ucsc.edu","firstName":"Jordan D.","lastName":"Ward","submittingAuthor":true,"correspondingAuthor":true,"equalContribution":null,"WBId":"","orcid":"0000-0001-9870-8936"}],"awards":[],"conflictsOfInterest":"The authors declare that there are no conflicts of interest present.
","dataTable":{"url":null},"extendedData":[],"funding":"This work was funded by the National Institutes of Health (NIH) National Institute of General Medical Sciences (NIGMS) (R01GM138701, and R35GM158317) to J.D.W.
","image":{"url":"https://portal.micropublication.org/uploads/5fca52711c3166238cb30a6745c2d5d9.png"},"imageCaption":"Tissue-specific RNAi strains used. A timed egg lay of animals of the indicated genotype were performed on control or mlt-11 (RNAi) plates and phenotypes were scored three days later. (B) Representative images of animals of the indicated genotype grown on control or mlt-11 RNAi. The red box highlights conditions that produced smaller larvae with molting defects. (C) Developmental delay was scored in the indicated strains grown on control or mlt-11 RNAi plates and classified as a failure to reach adulthood after 72 hours of growth. (D) Molting defects were scored in the indicated strains on control or mlt-11 RNAi. Scored defects included animals dragging cuticles, ecdysis failure and cuticle corsets. Tissue-specific RNAi data is from two independent replicates and the number of animals scored is in (A).
","imageTitle":"mlt-11 is necessary in seam cells for molting and larval development
","methods":"RNAi Knockdown
RNA interference experiments were performed as in Johnson et al. (2023). Control RNAi used an empty L4440. The mlt-11 (RNAi) vector was streaked from the Ahringer library (Kamath et al., 2003).
","reagents":"Strain name | Genotype | Source |
Bristol isolate, standard lab wild type | CGC | |
jsTi1493 {mosL loxP [wrdSi72(SCMp::pes-10delta::rde-1 CDS+3'UTR)] FRT3::mosR} IV ; rde-1(ne300) V | Ward lab Johnson et al., 2023) | |
jsTi1493 {mosL loxP [wrdSi97(suro-1p::rde-1 CDS+3'UTR)] FRT3::mosR} IV ; rde-1 (ne300) V | Ward lab Johnson et al., 2023) | |
CGC (Barkoulas et al., 2013) | ||
CGC (Melo & Ruvkun, 2012) | ||
CGC (Melo & Ruvkun, 2012) | ||
rde-1(ne300) V ; neIs9(neIs9 [myo-3::HA::RDE-1 + rol-6(su1006)]) X | CGC (Watts et al., 2020) | |
CGC (Espelt et al., 2005) |
Molting is an essential developmental process in nematodes, involving periodic synthesis of a new collagen-rich apical extracellular matrix (cuticle) and the shedding of the old cuticle. Understanding the mechanisms underlying molting can provide insight into oscillatory biological timers, epithelial biology, and is a source of potential targets to develop anthelminthics to control parasitic nematodes of medical and agricultural importance. Proteases and protease inhibitors play important and poorly understood roles in remodeling of the C. elegans aECM during molting.
mlt-11 is a putative Kunitz family protease inhibitor that plays a critical role in C.elegans cuticle synthesis and molting (Frand et al., 2005; Ragle et al., 2026). It is a complex protein with 10 Kunitz domains and depending on which of these domains are deleted, phenotypes such as left rollers, right rollers, microblistering, or embryonic lethality are observed (Ragle et al., 2026). Reduction of mlt-11 function results in aberrant protein localization in the aECM over both seam and hypodermal cells, and all three layers of the adult cuticle were affected (basal, medial, and cortical layers) (Ragle et al., 2026). Given that mlt-11 is expressed in both seam cells and in the hypodermal syncytium (Frand et al., 2005; Ragle et al., 2026), we turned to a tissue-specific RNAi approach to test where mlt-11 activity was needed to promote molting (Fig. 1A). mlt-11 knockdown in a tissue-specific RNAi strain (JDW371) that restricts knockdown to seam, hypodermal, and intestinal cells (Johnson et al., 2023), phenocopied mlt-11 RNAi in wild-type animals with respect to developmental delay and molting defects (Fig. 1B-D). mlt-11 RNAi in a hypodermal and seam cell-specific RNAi strain (QK52), produced less penetrant developmental delay and molting defects (Fig. 1B-D). Notably, mlt-11 RNAi in a hypodermal and intestinal-specific RNAi strain (JDW510), produced no developmental delay or molting defects, suggesting mlt-11 activity is necessary in seam cells and perhaps, in a limited capacity, in other hypodermal cells (Fig. 1B-D).
Despite robust hypodermal expression and localization (Frand et al., 2005; Ragle et al., 2026), our tissue-specific RNAi suggests that the seam cells are an important site of mlt-11 expression (Fig. 1). We consistently saw the most severe RNAi phenotypes in tissue-specific RNAi strains using a seam cell enriched promoter and no phenotypes from a hypodermal/intestinal-specific RNAi strain (Fig. 1B-D). Consistent with these data, we previously observed aberrant expression of the markers DPY-7::GFP (furrows), DPY-10::mScarlet (furrows), ROL-6::mNG (basal layer collagen), CUT-2::mNG (cortical layer cuticlin), and COL-19::mNG (basal layer collagen) over the seam cells as well as alae gaps following mlt-11 inactivation (Ragle et al., 2026). The medial layer strut collagen BLI-1 is normally excluded from the aECM over the seam cells but invades this area following mlt-11 knockdown (Ragle et al., 2026). These data were reminiscent of our work on NHR-23 as depletion of this factor causes aberrant aECM formation over the seam cells and nhr-23 activity also was necessary in the seam cells for developmental progression (Johnson et al., 2023). Seam cells are responsible for the production of alae (Singh & Sulston, 1978), so it is unclear why inactivation of nhr-23 or mlt-11 in the seam cells has such a potent effect on developmental progression and molting given that the hypodermis secretes the bulk of the cuticle. mlt-11 inactivation has been shown to cause reduced body length and width, impaired cuticle barrier function, and aberrant aECM protein localization (Ragle et al., 2026). Defining the tissue-specific requirements for mlt-11 in preventing these defects will be an important avenue for future investigation.
","references":[{"reference":"Barkoulas M, van Zon JS, Milloz J, van Oudenaarden A, Félix MA. 2013. Robustness and epistasis in the C. elegans vulval signaling network revealed by pathway dosage modulation. Dev Cell 24(1): 64-75.
","pubmedId":"23328399","doi":""},{"reference":"Espelt MV, Estevez AY, Yin X, Strange K. 2005. Oscillatory Ca2+ signaling in the isolated Caenorhabditis elegans intestine: role of the inositol-1,4,5-trisphosphate receptor and phospholipases C beta and gamma. J Gen Physiol 126(4): 379-92.
","pubmedId":"16186564","doi":""},{"reference":"Frand AR, Russel S, Ruvkun G. 2005. Functional genomic analysis of C. elegans molting. PLoS Biol 3(10): e312.
","pubmedId":"16122351","doi":""},{"reference":"Johnson LC, Vo AA, Clancy JC, Myles KM, Pooranachithra M, Aguilera J, et al., Ward JD. 2023. NHR-23 activity is necessary for C. elegans developmental progression and apical extracellular matrix structure and function. Development 150(10): 10.1242/dev.201085.
","pubmedId":"37129010","doi":""},{"reference":"Kamath RS, Fraser AG, Dong Y, Poulin G, Durbin R, Gotta M, et al., Ahringer J. 2003. Systematic functional analysis of the Caenorhabditis elegans genome using RNAi. Nature 421(6920): 231-7.
","pubmedId":"12529635","doi":""},{"reference":"Melo JA, Ruvkun G. 2012. Inactivation of Conserved C. elegans Genes Engages Pathogen- and Xenobiotic-Associated Defenses. Cell 149: 452-466.
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